Le T, Ferling I, Qin L, Nabaile C, Assuncao L, Roskelley CD, Grinstein S, and Freeman SA. Redistribution of the glycocalyx exposes phagocytic determinants on apoptotic cells. Developmental Cell, 2024

Chadwick S, Barreda D, Wu JZ, Ye G, Yusuf B, Ren D, and Freeman SA. Two pore channels regulate endomembrane tension to enable remodeling and resolution of phagolysosomes. PNAS, 2024

Barreda D, Grinstein S, and Freeman SA. Target lysis by cholesterol extraction is a rate limiting step in the resolution of phagolysosomes. European Journal of Cell Biology, 2024

Dissanayake D, Firouzabady A, Massumi M, de Paz Linares GA, Marshall C, Freeman SA, Laxer RM, Yeung RSM. Interleukin-1 mediated hyperinflammation in XIAP-deficiency is associated with defective autophagy. Blood, 2024

Lu C-Y, Wu JW, Yao HHY, Liu RJY, Li L, Pluthero FG, Freeman SA, Kahr WHA. Acidification of α-granules in megakaryocytes by vacuolar-type ATPase is essential for organelle biogenesis. Journal of Thrombosis and Haemostasis, 2024



Mylvaganam S and Freeman SA. The resolution of phagosomes. Immunological Reviews, 2023

de Paz Linares GA, Freeman SA, Cai R. Using Ion Substitution and Fluid Indicators to Monitor Macropinosome Dynamics in Live Cells. Phagocytosis and Phagosomes: Methods and Protocols, 2023

Wu J, Zeziulia M, Kwon W, Jentsch TJ, Grinstein S, Freeman SA. ClC-7 drives intraphagosomal chloride accumulation to support hydrolase activity and phagosome resolution. Journal of Cell Biology, 2023

Hirata Y, Cai R, Steinberg BE, Volchuk A, Saito Y, Matsuzawa A, Grinstein S, Freeman SA. Lipid peroxidation increases membrane tension, Piezo1 gating and cation permeability to execute ferroptosis. Current Biology, 2023

Teoh CW, Riedl Khursigara M, Ortiz-Sandoval CG, Park JW, Li J, Bohorquez-Hernandez A, Bruno V, Bowen EE, Freeman SA, Robinson LA, Licht C. The loss of glycocalyx integrity impairs complement factor H binding and contributes to cyclosporine-induced endothelial cell injury. Frontiers in Medicine, 2023

Freeman SA, Grinstein S, Orlowski J. Determinants, maintenance, and function of organellar pHPhysiological Reviews, 2023

Ghasempour S, and Freeman SA. The glycocalyx and immune evasion in cancer. The FEBS Journal, 2023



Yazdani S, Bilan PJ, Jaldin-Fincati JR, Pang J, Ceban F, Saran E, Brumell JH, Freeman SA, Klip A.  Dynamic glucose uptake, storage and release by human microvascular endothelial cells. MBoC, 2022

Mylvaganam S, Plumb J, Yusuf B, Li R, Lu CY, Robinson LA, Freeman SA, Grinstein S.  The spectrin cytoskeleton integrates endothelial mechanoresponses. Nature Cell Biology, 2022



Leung G, Zhou Y, Ostrowski P, Mylvaganam S, Boroumand P, Mulder DJ, Guo C, Muise AM, Freeman SA.  ARPC1B binds WASP to control actin polymerization and curtail tonic signaling in B cells. JCI Insight, 2021

Arif AA, Huang YH, Freeman SA, Atif J, Dean P, Lai JCY, Blanchet MR, Wiegand KC, McNagny KM, Underhill TM, Gold MR, Johnson P, Roskelley CD. Inflammation-induced metastatic colonization of the lung is facilitated by hepatocyte growth factor-secreting monocyte-derived macrophages.  Molecular Cancer Research, 2021

Saric A, Freeman SA, Williamson C, Jarnik M, Guardia C, Fernandopulle M, Gershlick D, Bonifacino J. SNX19 Restricts endolysosome motility through contacts with endoplasmic reticulumNature Communications 12(4552), 2021

Mylvaganam S, Freeman SA, Grinstein S. The cytoskeleton in phagocytosis and macropinocytosis. Current Biology 31(10), 619–632, 2021

Wang L, Aschenbrenner D, Zeng Z, Cao X, Mayr D, Mehta M, Capitani M, Warner N, Pan J, Wang L, Li Q, Zuo T, Cohen-Kedar S, Lu J, Ardy RC, Mulder DJ, Dissanayake D, Peng K, Huang Z, Li X, Wang Y, Wang X, Li S, Bullers S, Gammage AN, Warnatz K, Schiefer AI, Krivan G, Goda V, Kahr WHA, Lemaire M; Genomics England Research Consortium, Lu CY, Siddiqui I, Surette MG, Kotlarz D, Engelhardt KR, Griffin HR, Rottapel R, Decaluwe H, Laxer RM, Proietti M, Hambleton S, Elcombe S, Guo CH, Grimbacher B, Dotan I, Ng SC, Freeman SA, Snapper SB, Klein C, Boztug K, Huang Y, Li D, Uhlig HH, Muise AM. Gain-of-function variants in SYK cause immune dysregulation and systemic inflammation in humans and mice. Nature Genetics 53,4: 500-510, 2021

Chadwick SR, Grinstein S, Freeman SAFrom the inside out: Ion fluxes at the centre of endocytic traffic.  Current Opinion in Cell Biology 71, 77-86, 2021

Saric A and Freeman SA. Endomembrane Tension and TraffickingFrontiers in Cell and Developmental Biology. 8:611326, 2021

Saric A and Freeman SA. Solutes as controllers of endomembrane dynamicsNature Reviews Molecular Cell Biology, 2021

Chadwick SR, Wu JZ, Freeman SA  Solute Transport Controls Membrane Tension and Organellar Volume.  Cellular Physiology & Biochemistry. 55(S1):1-24, 2021

Freeman SA and Grinstein S. Promoters and Antagonists of Phagocytosis: A Plastic and Tunable Response Annual Review of Cell and Developmental Biology, volume 37, 2021



Kwon W and Freeman SAPhagocytosis by the Retinal Pigment Epithelium: Recognition, Resolution, Recycling.  Frontiers in Immunology. 11:604205, 2020

Imbert PRC, Saric A, Pedram K, Bertozzi CR, Grinstein S, Freeman SA. An Acquired and Endogenous Glycocalyx Forms a Bidirectional “Don’t Eat” and “Don’t Eat Me” Barrier to Phagocytosis. Current Biology, 2020

Westman J, Freeman SA, Grinstein S. Unconventional role of lysosomes in phagocytosis. Cell Calcium 91, 102269, 2020

Mercer J, Lee JE, Saphire EO, Freeman SASnapShot: Enveloped Virus Entry. Cell 182 (3), 786-786. e1, 2020

Mylvaganam S, Riedl M, Vega A, Collins RF, Jaqaman K, Grinstein S, Freeman SAStabilization of Endothelial Receptor Arrays by a Polarized Spectrin Cytoskeleton Facilitates Rolling and Adhesion of Leukocytes. Cell Reports 31 (12), 107798, 2020

Freeman SA, Uderhardt S, Saric A, Collins RF, Buckley CM, Mylvaganam S, Boroumand P, Plumb J, Germain RN, Ren D, Grinstein S. Lipid-gated monovalent ion fluxes regulate endocytic traffic and support immune surveillance. Science 367 (6475), 301-305, 2020

Freeman SA and Grinstein S. Phagocytosis: mechanosensing, traction forces, and a molecular clutch. Current Biology 30 (1), R24-R26, 2020



Marques PE, Nyegaard S, Collins RF, Troise F, Freeman SA, Trimble WS, Grinstein S. Multimerization and Retention of the Scavenger Receptor SR-B1 in the Plasma Membrane. Developmental Cell. 50 (3), 283-295, 2019

Ostrowski PP, Freeman SA, Fairn G, Grinstein S. Dynamic Podosome-Like Structures in Nascent Phagosomes Are Coordinated by Phosphoinositides. Developmental Cell. 50 (4), 397-410, 2019



Freeman SA and Grinstein S. Resolution of macropinosomes, phagosomes and autolysosomes: Osmotically driven shrinkage enables tubulation and vesiculation. Traffic. 19 (12), 965-974, 2018

Mylvaganam SM, Grinstein S, Freeman SAPicket-fences in the plasma membrane: functions in immune cells and phagocytosis. Semin Immunopathol. 40 (6), 605-615, 2018

Schlam D, Grinstein S, Freeman SA. Screening for Rho GTPase Modulators in Actin-Dependent Processes Exemplified by Phagocytosis. Methods Mol Biol. 107-127, 2018

Vega AR, Grinstein S, Freeman SA, Jaqaman K. Multistep Track Segmentation and Motion Classification for Transient Mobility Analysis. Biophysical Journal. 114(5): 1018-1025, 2018

Freeman SA, Vega AR, Riedl M, Collins RF, Ostrowski PP, Woods EC, Bertozzi CR, Tammi MI, Lidke DS, Johnson P, Mayor S, Jaqaman K, Grinstein S. Transmembrane pickets connect cyto-and pericellular skeletons forming barriers to receptor engagement. Cell. 172(1): 305-317, 2016



Marques PE, Grinstein S, Freeman SA. SnapShot: Macropinocytosis. Cell. 169(4): 766, 2017

Wang JC, Lee JY-J, Christian S, Dang-Lawson M, Pritchard C, Freeman SA, Gold MR. The Rap1–cofilin-1 pathway coordinates actin reorganization and MTOC polarization at the B cell immune synapse. Journal of Cell Science. 130(6): 1094-1109, 2017

Freeman SA, Christian S, Austin P, Iu I, Graves ML, Huang L, Tang S, Coombs D, Gold MR, Roskelley CD. Applied stretch initiates directional invasion through the action of Rap1 GTPase as a tension sensor. Journal of Cell Science. 130(1): 152-163, 2017

Sharfe N, Karanxha A, Dadi H, Merico D, Chitayat D, Herbrick J, Freeman SA, Grinstein S, Roifman CM. Dual loss of p110δ PI3-kinase and SKAP (KNSTRN) expression leads to combined immunodeficiency and multisystem syndromic features. Journal of Allergy and Clinical Immunology. 142(2): 618-629, 2017

Hua R, Cheng D, Coyaud É, Freeman SA, Di Pietro E, Wang Y, Vissa A, Yip CM, Fairn GD,  Braverman N, Brumell JH, Trimble WS, Raught B, Peter K Kim. VAPs and ACBD5 tether peroxisomes to the ER for peroxisome maintenance and lipid homeostasis. J Cell Biol. 216(8): 367-377, 2017



Saric A, Grinstein S, Freeman SAMeasurement of Autolysosomal pH by Dual-Wavelength Ratio Imaging. Methods in Enzymology. 588: 15-29, 2016

Freeman SA and Grinstein S. Phagocytosis: How Macrophages Tune Their Non-professional Counterparts. Current Biology. 26(24): 1279-1282, 2016

Wong HS, Jaumouillé V, Freeman SA, Doodnauth SA, Schlam D, Canton J, Mukovozov IM, Saric A, Grinstein S, Robinson LA. Chemokine signaling enhances CD36 responsiveness toward oxidized low-density lipoproteins and accelerates foam cell formation. Cell Reports. 14(12): 2859-71, 2016

Ostrowski PP, Grinstein S, Freeman SA. Diffusion barriers, mechanical forces, and the biophysics of phagocytosis. Developmental Cell. 38(2): 135-146, 2016

Oudhoff MJ, Braam MJS, Freeman SA, Wong D, Rattray DG, Wang J, Antignano F, Snyder K, Refaeli I, Hughes MR, McNagny KM, Gold MR, Arrowsmith CH, Sato T, Rossi FMV, Tatlock JH, Owen DR, Brown PJ, Zaph C. SETD7 controls intestinal regeneration and tumorigenesis by regulating Wnt/β-catenin and hippo/YAP signaling. Developmental Cell. 37(1): 47-57, 2016

Schlam D, Canton J, Carreño M, Kopinski H, Freeman SA, Grinstein S, Fairn GD. Gliotoxin Suppresses Macrophage Immune Function by Subverting Phosphatidylinositol 3, 4, 5-Trisphosphate Homeostasis. MBio. 7(2): 02242-15, 2016

Freeman SA, Goyette J, Furuya W, Woods EC, Bertozzi CR, Bergmeier W, Hinz B, van der Merwe PA, Das R, Grinstein S. Integrins form an expanding diffusional barrier that coordinates phagocytosis. Cell. 164(1): 128-140, 2016



Freeman SA, Jaumouillé V, Choi K, Hsu BE, Wong HS, Abraham L, Graves ML, Coombs D, Roskelley CD, Das R, Grinstein S, Gold MR. Toll-like receptor ligands sensitize B-cell receptor signalling by reducing actin-dependent spatial confinement of the receptor. Nature Communications. 6: 6168, 2015

Schlam D, Bagshaw R, Freeman SA, Collins RF, Fairn G, Grinstein S. Phosphoinositide 3-kinase enables phagocytosis of large particles by terminating actin assembly through Rac/Cdc42 GTPase-activating proteins. Nature Communications. 6: 8623, 2015


– 2014

Barsyte-Lovejoy, Li F, Oudhoff MJ, Tatlock JH, Dong A, Zeng H, Wu H, Freeman SA, Schapira M, Senisterra GA, Kuznetsova E, Marcellus R, Allali-Hassani A, Kennedy S, Lambert JP, Couzens AL, Aman A, Gingras AC, Al-Awar R, Fish PV, Gerstenberger BS, Roberts L, Benn CL, Grimley RL, Braam MJ, Rossi FM, Sudol M, Brown PJ, Bunnage ME, Owen DR, Zaph C, Vedadi M, Arrowsmith CH. (R)-PFI-2 is a potent and selective inhibitor of SETD7 methyltransferase activity in cells. Proceedings of the National Academy of Sciences. 111(35): 12853-12858, 2014

Freeman SA and Grinstein S. Phagocytosis: receptors, signal integration, and the cytoskeleton. Immunological Reviews. 262(1): 193-215, 2014

Oudhoff MJ, Freeman SA, Couzens AL, Antignano F, Kuznetsova E, Min PH, Northrop JP, Lehnertz B, Barsyte-Lovejoy D, Vedadi M, Arrowsmith CH, Nishina H, Gold MR, Rossi FMV, Gingras A, Zaph C. Control of the hippo pathway by set7-dependent methylation of yap. Developmental Cell. 26(2): 188-194, 2013

Austin P, Freeman SA, Gray CA, Gold MR, Vogl AW, Andersen RJ, Roberge M, Roskelley CD. The invasion inhibitor sarasinoside A1 reverses mesenchymal tumor transformation in an E-cadherin-independent manner. Molecular Cancer Research. 11(5): 530-40, 2013

Maltby S, Freeman SA, Gold MJ, Baker JHE, Minchinton AI, Gold MR, Roskelley CD, McNagny KM. Opposing Roles for CD34 in B16 Melanoma Tumor Growth Alter Early Stage Vasculature and Late Stage Immune Cell Infiltration. PloS one. 6(4): 18160, 2011

Freeman SA, Lei V, Dang-Lawson M, Mizuno K, Roskelley CD, Gold MR. Cofilin-Mediated F-Actin Severing Is Regulated by the Rap GTPase and Controls the Cytoskeletal Dynamics That Drive Lymphocyte Spreading and BCR Microcluster Formation. The Journal of Immunology. 187(11): 5887-5900, 2011

Davies AH, Barrett I, Pambid MR, Hu K, Stratford AL, Freeman SA, Berquin IM, Pelech S, Hieter P, Maxwell C, Dunn SE. YB-1 evokes susceptibility to cancer through cytokinesis failure, mitotic dysfunction and HER2 amplification. Oncogene. 30(34): 3649-60, 2011

Freeman SA, McLeod SJ, Dukowski J, Austin P, Lee CCY, Millen-Martin B, Kubes P, McCafferty D, Gold MR, Roskelley CD. Preventing the activation or cycling of the Rap1 GTPase alters adhesion and cytoskeletal dynamics and blocks metastatic melanoma cell extravasation into the lungs. Cancer Research. 70(11): 4590-601, 2010

Lin KBL, Freeman SA, Gold MR. Rap GTPase-mediated adhesion and migration: A target for limiting the dissemination of B-cell lymphomas?. Cell adhesion & migration. 4(3):327-32, 2010

Lin KBL, Tan P, Freeman SA, Lam M, McNagny KM, Gold MR. The Rap GTPases regulate the migration, invasiveness and in vivo dissemination of B-cell lymphomas. Oncogene. 29(4): 608-615, 2009

Lin KBL, Freeman SA, Zabetian S, Brugger H, Weber M, Lei V, Dang-Lawson M, Kathy WK,  Santamaria R, Batista FD, Gold MR. The rap GTPases regulate B cell morphology, immune-synapse formation, and signaling by particulate B cell receptor ligands.  Immunity. 28(1): 75-87, 2008

Badour K, McGavin MKH, Zhang J, Freeman SA, Vieira C, Filipp D, Julius M, Mills GB, Siminovitch KA. Interaction of the Wiskott-Aldrich syndrome protein with sorting nexin 9 is required for CD28 endocytosis and cosignaling in T cells. Proceedings of the National Academy of Sciences. 104(5): 1593-1598, 2007

Leng Y, Zhang J, Badour K, Arpaia E, Freeman SA, Cheung P, Siu M, Siminovitch K. Abelson-interactor-1 promotes WAVE2 membrane translocation and Abelson-mediated tyrosine phosphorylation required for WAVE2 activation.  Proceedings of the National Academy of Sciences. 102(4): 1098-1103, 2005